棘颊雀鲷人工繁殖与育苗的初步研究

为了解棘颊雀鲷(Premnas biaculeatus)在人工饲养条件下繁殖和仔鱼培养对光照、水温及饵料的要求,采用实验生物学的方法,在严格控制的条件下培养亲鱼、胚胎和仔鱼.结果表明,在(26±1)℃和14L:10D的光暗周期下,2 000 1x和500 1x两种光照强度对亲鱼产卵周期和卵的受精率并没有明显的影响,但是...     

Spatial structures of hydrothermal vents and vent-associated megafauna in the back-arc basin system of the Okinawa Trough,western Pacific

Spatial patterns and morphology of hydrothermal vents and the occurrence of vent-associated megafauna were investigated in the back-arc basin system of the Okinawa Trough, western Pacific. Amongst hydrothermal vent fields located on the rising slopes to the NE and S of the basin, the Iheya North area has been subjected to a series of intensive diving surveys. Hydrothermal vents demonstrated concentrated patterns of distribution on different spatial scales. In Iheya North, the majority of vents occurred in lines parallel to the NE to W/SW alignment of the spreading axis, within a radius of ca. 200 m around the most active sulphide structure with the highest recorded fluid temperature of over 300°C. The morphology of hydrothermal vents varied greatly from an incipient flat rock with crevices to a 20- to 30-m-tall, multi-flanged structure with concomitant variation in the distribution and abundance of vent-associated megafauna, particularly a galatheid Shinkaia crosnieri and Bathymodiolus mussels. Comparison of active and inactive vents revealed that the spatial extent of Shinkaia ‘aggregation’ (a group of individuals with short nearest-neighbour distances) effectively defined a habitat unit of this species, and active hydrothermal chimneys contained more of these units, leading to greater spatial occupancy by Shinkaia. Given the temporally unstable nature of vent structures as recognised by repeated surveys, vent assemblages are thought to be closely dictated by the spatio-temporal dynamics of vents in the Okinawa Trough back-arc system.     

Colonization of Sulfurovum sp. on the gill surfaces of Alvinocaris longirostris, a deep-sea hydrothermal vent shrimp

Deep‐sea hydrothermal vents are unique light‐independent ecosystems that are sustained by chemosynthetic bacteria. For many of the invertebrates inhabiting in such environments, bacteria play essential roles in both energy acquisition and detoxification of potentially toxic gases such as H₂S. In this study, the bacterial flora present on the gills of Alvinocaris longirostris (Bresiliidae: Caridea), a shrimp inhabiting hydrothermal vents (1532 m depth) at the Hatoma Knoll of the Okinawa Trough, was investigated. Bacterial 16S rDNA fragments were successfully amplified from the gills and 70% of these fragments showed an identical pattern in the restriction fragment length polymorphism analysis. These fragments were assigned to the ribotype AL‐1. Phylogenetic analyses suggest that AL‐1 forms a monophyletic clade with Sulfurovum spp. (ε‐Proteobacteria). Fluorescence in situ hybridization for AL‐1 and electron microscopy showed the presence of short‐rod bacteria lining up on the cuticular layer of the surface of the gill filaments. These results suggest that bacterial association with gills also occurs in bresiliid shrimps.     

Dispersion of a tracer on the East Pacific Rise (9°N to 10°N), including the influence of hydrothermal plumes

On 12 November 2006, 3 kg of sulfur hexafluoride were released in a 1.2 km long streak in the axial summit trough of the East Pacific Rise at 9°30′N to study how circulation and mixing affect larval dispersion. The first half of a tracer survey performed approximately 40 days after the injection found a small percentage of the tracer on the ridge axis between 9°30′N and 10°10′N, with the main concentration near 9°50′N, a site of many active hydrothermal vents. These observations provide evidence of larval connectivity between vent sites on the ridge. The latter half of the survey detected the primary patch of tracer west of the ridge and just south of the Lamont Seamounts, as a majority of the tracer had been transported off the ridge. However, by the end of the survey, the eastern edge of this patch was transported back to within 10 km of the ridge crest at 9°50′N by a reversal in the subinertial flow, suggesting another pathway for larvae between points along the ridge. Both the horizontal and vertical distributions of the tracer were complex and were likely heavily influenced by topography and vents in the area. Elevated tracer concentrations within the axial summit trough and an adjacent depression on the upper ridge flank suggest that tracers may be detained in such depressions. Correlated tracer/turbidity profiles provide direct evidence of entrainment of the tracer into vent plumes from 9°30′N to 10°N. A comparison of the vertical tracer inventory with neutral density vent-plume observations suggests that on the order of 10% of the tracer injected was entrained into vent plumes near the injection site. The results imply that effluent from diffuse hydrothermal sources and larvae of hydrothermal vent fauna can be entrained in significant quantities into plumes from discrete sources and dispersed in the neutrally buoyant plumes.     

Common bivalve larvae from New Zealand: Leptonacea

The late stage larvae of three erycinid bivalves (Mollusca: Pelecypoda: Leptonacea) taken from the brood chamber of the adult are described (Kellia cycladiformis, Lasaea rubra hinemoa, and L. maoria), and the provisionally identified late stage larva of the erycinid Arthritica bijurca taken from the plankton is described. Also, the D‐shaped larvae of the erycinid bivalves Kellia cycladiformis, Borniola reniformis, Arlhritica crassijormis, and A. bifurca taken from the parent brood chamber are described. The seasonal occurrence of each late stage larva in the plankton at the Bay of Islands (35° 15'S, 174° 10'E), Wellington Harbour (41° 16'S, 174° 51'E), and Raumati Beach (40° 56'S, 174° 58'E), New Zealand is described. Aspects of the reproductive cycles of Lasaea rubra hinemoa and Arthritica bifurca are presented.     

Benthic fauna associated with mussel beds and shrimp swarms at hydrothermal fields on the Mid-Atlantic Ridge

Macrofaunal assemblages with the prevalence of Bresiliidae shrimp and Mytilidae mussels are abundant in the hydrothermal vents along the Mid-Atlantic Ridge. The mussels inhabit the zone of diffuse seeps of hydrothermal fluids with temperature abnormalities up to several degrees. Shrimps inhabit an extreme biotope in the mixed interface between the seawater and the hydrothermal fluid at a temperature up to 20–30°C. We studied the mussel and shrimp assemblages in three hydrothermal vent fields: the Rainbow, Broken Spur, and Snake Pit. The species richness of the mussel assemblages in at least two regions (Broken Spur and Snake Pit) is higher as compared with the shrimps of the same hydrothermal vent fields. The fauna inhibiting the shrimp swarms lack almost any taxa specific for particular assemblages: almost all the taxa are also present in the mussel beds. The structure of the shrimp assemblage is less homogeneous as compared with that of the mussel assemblage. The population prevalence of one taxon (Copepoda) in the shrimp assemblage is most likely connected with the extreme and unstable conditions of the biotope occupied by the shrimps in the hydrothermal field. The taxonomic similarity between the mussel and shrimp assemblages within one hydrothermal vent field is higher as compared with the similarity between the mussel (or shrimp) assemblages from different fields.     

环境温度和盐度对墨吉明对虾(Fenneropenaeus merguiensis)胚胎发育的影响

以自然成熟的亲虾为材料,观察并描述了墨吉明对虾(Fenneropenaeus merguiensis)从受精卵到幼体孵化出膜的胚胎发育过程。结果显示,在水温29°C,盐度28的条件下,墨吉明对虾整个胚胎发育过程需要11h左右。水温和盐度对幼体出膜时间和受精卵的孵化率均有显著的影响(P0.05)。孵化水温从32°C下降到23°C时,幼体出膜时间由555min(9.25h)增加到1108min(18.46h)。胚胎发育的最适水温为26—32°C,在此温度范围内,墨吉明对虾的幼体出膜时间与水温呈负相关。胚胎发育的最适盐度25—35,在此范围内,墨吉明对虾的幼体出膜时间和孵化率均无显著差异(P0.05),说明墨吉明对虾的胚胎发育的盐度范围较广。     

Early development of Forsterygion varium,Gilloblennius decemdigitatus,and G. tripennis (Pisces: Tripterygiidae)

In the Wellington area (central New Zealand) Forsterygion varium (Forster, in Bloch & Schneider 1801) spawns from early June to late November, Gilloblennius decemdigitatus (Clarke, 1879) from early June to early October, and G. tripennis (Forster, in Bloch & Schneider 1801) from July to October. They deposit their eggs on firm substrates below the low tide level. Egg clusters are attended by an adult fish until hatching, which in the laboratory at 11.5–13.5°C, occurs after 18 days for F. varium and 20 days for G. decemdigitatus. Development to hatching for G. tripennis takes about 21 days at 13–15°c. The egg development of all three species is illustrated. The yolk‐sac larvae (prolarvae) of F. varium, G. decemdigitatus, and G. tripennis at hatching average 5.85, 5.03, and 5.72 mm standard length respectively. Yolk‐sac larvae of F. varium have a single row of 9–17 small, stellate melanophores along the ventral midline of the tail. The yolk‐sac larvae of G. tripennis have up to nine melanophores along the ventral midline of the tail, a medium‐sized melanophore above the optic lobes, and a further two above the medulla. The yolk‐sac larvae of G. decemdigitatus have three large mid‐dorsal melanophores interspersed with five smaller, paler pigment cells, and three large mid‐ventral melanophores interspersed with three of the smaller cells.     

Carapidae larvae (Acanthomorpha,Paracanthopterygii, Ophidiiformes) from the oceanic region of the southwest Atlantic Ocean off Brazil (12–22°S)

We report the occurrence and distribution of Carapidae larvae in the southwest Atlantic Ocean, between Real River (12°S) and São Tomé Cape (22°S), off Brazil. Carapidae larvae were collected during three oceanographic cruises: spring 1998, winter 1999 and autumn 2000. The hauls were conducted obliquely from a maximum depth of 200 m, during day and night, with bongo nets 330- and 500-μm mesh size. Three species were recorded in this study: Carapus bermudensis, Echiodon dawsoni and Snyderidia canina. Only one C. bermudensis was collected during the winter cruise off the Salvador coast (13°S). Twenty-one larvae of E. dawsoni were collected during the three cruises between 14.5° and 22°S, including some at the banks of Vitória–Trindade Ridge. One S. canina larva occurred, during the spring in the north coast of Rio de Janeiro (22°S).     

Latitudinal variation in growth of Crangon crangon (L.): Does counter-gradient growth compensation occur?

Length growth in relation to water temperature was studied for Crangon crangon (L.) from two populations at the northern and southern edges of its distributional range to determine whether counter-gradient growth compensation occurs. In crustaceans, growth rate depends on the time between moulting events (intermoult period) and the size increase at moult (moult increment). In this study, the period between moults was shorter at higher temperature, ranging respectively from about 11 days at 25 °C to 27 days at 10 °C at southern edge, and from 10 to 24 days at the same temperatures at the northern edge. Moult increment showed a large variability, from 1.5 to 2.7 mm with no clear trend with temperature at the northern edge; and decreasing from about 2.7 mm at 10 °C to about 1.5 mm at 25 °C at the southern edge. As a result, the temperature effect on the overall growth rate differed between shrimps from the north and those from the south, suggesting counter-gradient growth compensation. At the northern edge, mean growth increased from about 0.12 mm d^− 1 at 10 °C to about 0.23 mm d^− 1 at 25 °C, while at the southern edge, growth was lower, about 0.08 mm d^− 1 at 10 °C and increased to about 0.16 mm d^− 1 at 25 °C. Maximum observed growth rates of shrimps from the north were also higher and ranged from 0.17 mm d^− 1 at 10 °C to 0.89 mm d^− 1 at 25 °C, while shrimps from the south grew at a maximum of 0.08 to 0.75 mm d^− 1 respectively at 15 and 20 °C. Sex and size differences were also found, with males growing slower than females and at a decreasing growth rate with increasing size. Implications for the brown shrimp's life cycle are discussed.     

Modeled interactions of mesoscale eddies with the East Pacific Rise: Implications for larval dispersal

Larval transport from distant populations is essential for maintenance and renewal of populations in patchy and disturbed ecosystems such as deep-sea hydrothermal vents. We use quasi-geostrophic modeling to consider the potential for long-distance dispersal of hydrothermal vent larvae in mesoscale eddies interacting with the northern East Pacific Rise. Modeled eddy dynamics were similar to the observed propagation dynamics of Tehuantepec eddies, including their ability to cross the ridge. Simulated surface anticyclones were associated with coherent cyclones in the deep layer with relatively strong current velocities that could significantly increase the dispersal potential of passive particles. Eddy interactions with ridge topography further enhanced tracer dispersal along the ridge axis through shearing and elongation of the eddy core. Simulations suggest that the passage of an eddy would result in local loss from the vent field and aggregate transport with potential enhancement of dispersal between vent fields separated by up to 270 km. Based on the latitude at which most Tehuantepec eddies cross the ridge, eddy-induced flows would enhance connectivity between the 13°N, 11°N, and 9°N vent fields along the East Pacific Rise asymmetrically with higher transport from northern vent fields to southern vent fields.     

Community structure in mussel beds at Logatchev hydrothermal vents and a comparison of macrofaunal species richness on slow- and fast-spreading mid-ocean ridges

Species lists for vent fields on the Mid‐Atlantic Ridge (MAR) from 14°N to 38°N suggest that there is a northern (>27°N), shallow (<2000 m) fauna and a southern (<27°N), deeper (>3000 m) endemic vent fauna, but little is known about how community structure varies along the ridge axis. In this study, quantitative samples of macrofaunal invertebrates associated with mussels (Bathymodiolus puteoserpentis) were collected at Logatchev (14°45′N), the southern‐most explored vent field on the MAR. Community structure (including species composition, species richness, diversity, and relative species abundances) in mussel beds at Logatchev was compared with that of Snake Pit (23°22′N) and Lucky Strike (37°17′N) mussel beds. The most striking feature of the Logatchev mussel‐bed macrofaunal invertebrate community was the tremendous abundance (up to 2390 individuals per liter of mussel‐volume sampled) and biomass of the ophiuroid, Ophioctenella acies. Logatchev and Snake Pit mussel beds share >50% of their associated macrofaunal species; these two sites share only 20–25% of their macrofaunal species with Lucky Strike. Species–effort curves and univariate measures of diversity (H′, J′) do not support the claim that diversity of vent organisms on the MAR is highest at Logatchev, at least when one assesses this within a habitat type. Multivariate analysis readily differentiates the species‐abundance characteristics of Logatchev, Snake Pit, and Lucky Strike mussel‐bed macrofaunas. The relationship between sea‐floor spreading rate and diversity was explored through comparison of species richness in mussel‐bed habitats on slow‐spreading (MAR), fast‐spreading [northern East Pacific Rise (EPR)], and ultra‐fast‐spreading (southern EPR) mid‐ocean ridges. Species richness was greater in samples from the faster‐spreading ridge axes, where vents are more closely spaced but shorter lived, than on slow‐spreading centers, where vents are further apart but longer lived.     

Molecular diversity of Thermococcales isolated from Guaymas Basin hydrothermal vents

Members of Thermococcales are organoheterotrophic hyperthermophilic anaerobes consisting of three genera, Thermococcus, Pyrococcus, and Palaeococcus, among which Thermococcus are the most frequently isolated. In this study, a variety of Thermococcales strains were isolated from the Guaymas Basin hydrothermal vents under different temperatures and pressures. Based on their 16S rRNA gene sequences, all the strains isolated from 85 C or 95 C enrichment cultures at ambient pressure were classified as Thermococcus, while strains isolated from 108 C-30 MPa enrichment cultures belonged to Pyrococcus. The combination of high pressure and high temperature for enrichment was shown to be an efficient method for isolating Pyrococcus strains. Through selecting and analyzing with multiple molecular phylogenetic markers, the isolated Thermococcales strains were found quite diversified including several putative novel species. This demonstrates a vast genetic reservoir of Thermoccoccales in the deep-sea hydrothermal vents, and a fast evolving of the Thermococcales species adapting to the fast changing environment.     

Temperature effects on the embryo development and hatching of the spiny lobster Sagmariasus verreauxi

Factors affecting the development of the embryos and production of larvae of the spiny lobster Sagmariasus verreauxi were investigated. Mature lobsters were collected from the fishery in New Zealand and held in captivity until they moulted, mated, and extruded eggs. For females, moulting took place in winter (July/August) and mating occurred 58–88 days later in early spring. The ovigerous females were then held at three temperatures and embryo development monitored at approximately weekly intervals. Mean times to hatch were 55, 75, and 130 days at 20°C, 17°C, and 13°C respectively. The total number of larvae that hatched from each female ranged from 0.97 to 1.4 million. The theoretical temperature at which embryo development ceases (a biological zero) for S. verreauxi, of 9.4°C, was calculated from data on the appearance of the median eye, the eyes and the chromatophores, and the time to hatch. An eye index formula was also derived to allow prediction of time to hatch at a range of temperatures using the cumulative difference between the rearing temperature and the biological zero. First instar phyllosomas from embryos reared at 20°C were significantly smaller than those from 17°C and 13°C but there was no significant difference in fitness. This study shows that hatching can be spread over a number of months, without significantly affecting larval quality, simply by manipulating holding temperature.     

Ecophysiological differences between vesicomyid species and metabolic capabilities of their symbionts influence distribution patterns of the deep‐sea clams

This study provides an analysis of vesicomyid bivalve–symbiont community distribution across cold seep and hydrothermal vent areas in the Guaymas Basin (Gulf of California, Mexico). Using a combination of morphological and molecular approaches including fluorescent in situ hybridization (FISH), and electronic microscopy observations, vesicomyid clam species and their associated symbionts were characterized and results were analyzed in light of geochemical conditions and other on‐site observations. A greater diversity of vesicomyids was found at cold seep areas, where three different species were present (Phreagena soyoae [syn. kilmeri], Archivesica gigas, and Calyptogena pacifica). In contrast, A. gigas was the only species sampled across the hydrothermal vent area. The same haplotype of A. gigas was found in both hydrothermal vent and cold seep areas, highlighting possible contemporary exchanges among neighboring vents and seeps. In either ecosystem, molecular characterization of the symbionts confirmed the specificity between symbionts and hosts and supported the hypothesis of a predominantly vertical transmission. In addition, patterns of clams could reflect potential niche preferences for each species. The occurrence of numerous traces of vesicomyid movements on sediments in the sites colonized by A. gigas seemed to indicate that this species might have a better ability to move. Furthermore, variation in gill sulfur content could reveal a higher plasticity and sulfur storage capacity in A. gigas. Thus, the distribution of vesicomyid species across the chemosynthetic areas of the Guaymas Basin could be explained by differences in biological traits of the vesicomyid species that would allow A. gigas to more easily exploit transient and punctual sources of available sulfide than P. soyoae.     

Population biology of Excirolana armata (Dana, 1853) (Isopoda,Cirolanidae) on an exposed sandy beach in Southeastern Brazil

The reproductive biology and population dynamics of the cirolanid isopod Excirolana armata (Dana, 1853) were analysed through monthly samples from December 2003 to November 2005 on Una beach, São Paulo state (24° S), in Southeastern Brazil. Sampling was performed along three transects established from the base of foredunes to the waterline. On Una beach, E. armata showed continuous reproduction with higher abundances of ovigerous females in winter and spring (July–November) with a higher peak of juveniles in spring (November 2004). The fecundity ranged from 2 to 18 eggs/embryos per female, depending on the female length. The incubation period was estimated as 2 months. The life span of males and females was nearly 1 year. The short life span and the high energetic expenditure inherent to reproduction with maternal care, probably kept females from producing more than one brood in their lifetime. When comparing the population of E. armata on Una beach (24° S) with populations in Southern Brazil (32° S), Uruguay (34° S) and Argentina (36° S), it was verified that several biological population traits (length of the smallest juvenile, length of the largest individual, length of the smallest and largest ovigerous females, range of fecundity and life span) tended to increase at higher latitudes, whereas other traits (instantaneous rate of mortality and the curvature parameter of von Bertalanffy growth function) tended to decrease. However, comparing E. armata on Una beach (24° S) with a population situated at a close latitude (25° S), unexpected differences in relation to population structure and to growth demonstrated and reinforced the importance of density‐dependent factors over life history traits of E. armata on dissipative beaches.     

Geologic setting of the Snake Pit hydrothermal site: An active vent field on the Mid-Atlantic Ridge

TheSnake Pit Hydrothermal Site lies on the axis of the Mid-Atlantic Ridge at 23°22′ N latitude, about 30 km south of the Kane Transform Intersection. Active ‘black smoker’ vents and a surrounding field of hydrothermal sediment occur at the crest of a laterally extensive neovolcanic ridge. It is one of the first active hydrothermal vent fields to be found on a slow-spreading ridge axis and despite significant differences in its geologic setting from those of the East Pacific Rise, has many similarities to its fast-spreading counterparts. Although preliminary reports have documented many interesting aspects of these vents and their surroundings, new data collected from the manned submersible ALVIN and the deep-towed ANGUS camera system define the regional tectonic setting as well as the local geologic environment of this fascinating area. The Snake Pit vents are located on a local peak of a volcanic constructional ridge at a depth of 3450 m, 700–800 m deeper than vents known from the East Pacific Rise, Galapagos, or Juan de Fuca spreading centers. The vent field is at least 600 m long and up to 200 m wide and is covered by a thick blanket of greenish to yellow-orange hydrothermal sediment. Both active and extinct vents are perched along the crests of steep-sided sulfide mounds that reach heights of over 40 m. High-temperature (350° C) fluids are vented from black smoker chimneys and low-temperature (226° C) fluids seep from sulphide domes and subordinate anhydrite constructions. Water temperatures, flow rates, fluid chemistries, and mineralization are strikingly similar to vents of faster spreading ridge crests; however, a somewhat distinct fauna inhabit the area.     

The effect of temperature and salinity on survival and growth of Crangon uritai (Decapoda: Crangonidae) larvae reared in the laboratory

Larvae of the decapod Crangon uritai were reared in the laboratory in a factorial experiment employing three temperatures (9, 12 and 15 °C) and three salinities (29‰, 32‰ and 35‰) from hatching to the post‐larval stage. The effects of temperature and salinity on survival, intermolt period (IP) and molt increment (MI) were investigated. Larvae from one brood were subdivided into groups of 20 and reared in glass bowls containing filtered sea‐water at a number of temperature–salinity combinations. The reared larvae were transferred daily to the clean bowls prepared with newly hatched Artemianauplii, and number of molts and mortality within each bowl were recorded. The zoeal size (carapace length) was determined from exuvia and dead larvae, and the IP was also recorded. Larvae of C. uritai completed larval development only at 15 °C temperature. The first zoeal stage completed their development at all temperature–salinity combinations and exhibited the highest survival rate. IP at each stage increased with increasing size and greatly decreased with increasing temperature, and intermolt duration (range of days) increased with larval development especially at the lower temperature. Although the MI decreased with increasing size, it was little affected by temperature. This led to a better growth rate with increasing temperature.     

Geochemistry and stable isotope constraints on high-temperature activity from sediment cores of the Saldanha hydrothermal field

The Saldanha hydrothermal field is hosted atop a mafic–ultramafic seamount, located at a non-transform offset on the Mid-Atlantic Ridge. Previous observations revealed a field where transparent low-temperature fluids discharge through centimeter-sized vents without the formation of chimney structures. We present geochemical and stable isotope (O and C) analyses from sediment samples collected at this field, both at and far from the vent area. Most sediments, including some directly adjacent to orifice vents, are pelagic oozes with only a weak hydrothermal overprinting. Hydrothermal precipitates are characterized by Fe–Mn oxyhydroxides and a minor amount of Cu–Zn sulphide minerals. However, one of the cores (SCD7) collected at the vent area shows a much stronger hydrothermal signature. This core is composed of a matrix of serpentine + talc ± chlorite with high porosity, where calcite + chalcopyrite + sphalerite/wurtzite ± pyrite–pyrrhotite were precipitated. In this core, metal enrichments, REE patterns, and the oxygen and carbon isotope composition of calcites indicate that mineralization must have occurred in the subsurface by high-temperature fluids, with minor mixing with seawater and with a significant magmatic contribution. Thus, while most samples confirm previous findings indicating that Saldanha hydrothermal fluid discharge is mainly diffuse and of low temperature, data from core SCD7 suggest that areas of high-temperature hydrothermal activity also occur, where temperatures of the fluids could reach > 260 °C and maximum temperatures of 330 °C. We suggest that fluids can flow through faults at the top of the mount and discharge in a more focused way through vent orifices, producing intense hydrothermal alteration of the sediments. At these locations complex hydrothermal processes occur, including reactions of the hydrothermal fluids with mafic and ultramafic rocks and magma degassing, as suggested by the carbon isotope composition of hydrothermal calcites. The high temperature of the fluid inferred from the geochemistry of the hydrothermal minerals requires a significant heat input to the system, suggesting an additional magmatic heat source to the already proposed exothermic serpentinization reactions.